Ecological Indicators

The suitable ecological water level of the East Dongting Lake based on the habitat
suitability of migratory birds feeding
--Manuscript Draft--

Manuscript Number: ECOLIND-20748

Article Type: Research paper

Keywords: Three Gorge Dam; ecological water level; habitat suitability; land cover; Dongting
Lake Wetland

Abstract: Understanding responses of ecological indicators to the water level is significant for
providing countermeasures of wetland protection. East Dongting Lake serves critical
wintering habitat for migratory birds along the East Asian-Australasian Flyway.
However, the mechanistic relationships between the water level and the suitable
habitat area of migratory birds feeding (WUA ) remain poorly characterized.
Therefore, we established a habitat suitability model of migratory birds feeding based
on DEM and NDVI, and recognized the change of WUA by comparing the transition of
land-cover of similar hydrologic years before and after the operation of Three Gorge
Dam (TGD). The results demonstrated that (1) Based on the habitat suitability model
of migratory birds feeding, the most suitable ecological water level in East Dongting
Lake is 20.47-21.55 m. (2) Compared with the pre-TGD period, the most prominent
alteration of WUA occurred in January, which increased by 6.51%. (3) The most
prominent transition among changes of land-cover were that the Reed beach (RB)
(155.07 km 2 ) and Marsh land (ML) (150.66 km 2 ) changed to Tender Marsh land
(TML) during dry hydrologic years, and the RB (171.90 km 2 ) and ML (174.30 km 2
) changed to TML during normal hydrologic years. (4) The increase of the water level in
dry seasons and duration days of water level from 28 to 30 m in wet seasons were the
main reasons for the improvement of habitat suitability of migratory birds. The findings
can provide a useful model for habitat conservation of migratory birds and a reference
for the operation of TGD.

Powered by Editorial Manager® and ProduXion Manager® from Aries Systems Corporation
Highlights (for review)

A habitat suitability model of migratory birds feeding based on DEM and NDVI.

Transition of land-cover of similar hydrologic years before and after 2003.

The driving factors for improving habitat suitability of migratory birds feeding.
Manuscript Click here to view linked References

1 The suitable ecological water level of the East Dongting Lake based

2 on the habitat suitability of migratory birds feeding

3 Yongwei Zhu, Hongxiang Wang, Wenxian Guo*

4 North China University of Water Resources and Electric Power, Zhengzhou 450045, China

5 ∗ Corresponding author: guowenxian163@163.com (W. Guo), Tel: +86-150-37121238

7 Abstract: Understanding responses of ecological indicators to the water level is significant for

8 providing countermeasures of wetland protection. East Dongting Lake serves critical wintering

9 habitat for migratory birds along the East Asian-Australasian Flyway. However, the mechanistic

10 relationships between the water level and the suitable habitat area of migratory birds feeding (WUA)

11 remain poorly characterized. Therefore, we established a habitat suitability model of migratory birds

12 feeding based on DEM and NDVI, and recognized the change of WUA by comparing the transition

13 of land-cover of similar hydrologic years before and after the operation of Three Gorge Dam (TGD).

14 The results demonstrated that (1) Based on the habitat suitability model of migratory birds feeding,

15 the most suitable ecological water level in East Dongting Lake is 20.47-21.55 m. (2) Compared with

16 the pre-TGD period, the most prominent alteration of WUA occurred in January, which increased

17 by 6.51%. (3) The most prominent transition among changes of land-cover was that the Reed beach

18 (RB) (155.07 km2) and Marsh land (ML) (150.66 km2) changed to Tender Marsh land (TML) during

19 dry hydrologic years, and the RB (171.90 km2) and ML (174.30 km2) changed to TML during

20 normal hydrologic years. (4) The increase of the water level in dry seasons and duration days of

21 water level from 28 to 30 m in wet seasons were the main reasons for the improvement of habitat

22 suitability of migratory birds. The findings can provide a useful model for habitat conservation of

23 migratory birds and a reference for the operation of TGD.

24 Keywords: Three Gorge Dam, ecological water level, habitat suitability, land cover, Dongting Lake

25 Wetland

26 1. Introduction
27 Wetlands, the cradles of biodiversity, play a significant role in water purification, flood storage

28 and drought resistance, and climate regulation (Sajedipour et al., 2017; Zhang et al., 2018a).

29 However, climate change and human activities had a remarkable impact on natural wetlands, which

30 leads to the change of wetland hydrologic regimes, water shortage, water quality deterioration, area

31 shrinkage, and function degradation (Wang et al., 2017; Zhou et al., 2019). Studies estimate that 54%

32 of the world’s wetlands have been lost since 1900 (Zhang et al., 2018a). The environmental and

33 ecological problems of wetlands have attracted extreme attention around the world (Stone, 2008;

34 Geng et al., 2021; Wu and Zheng, 2020; Zou et al., 2018). Researchers are looking for a scheme of

35 hydrologic regulation and water resources conservation for wetland ecological protection and

36 restoration (Zhang et al., 2016; Zhang et al., 2018a). In the above context, an increasing number of

37 studies began to focus on the Ecological Water Requirements (EWRs) of wetlands (Sajedipour et

38 al., 2017; Prakasam et al., 2021; Zhang et al., 2016).

39 The EWRs are water regimes needed to ensure species diversity and ecological integrity at a

40 low level of risk (Gleick, 1996). The methods of EWRs are mainly categorized into four groups

41 including hydrology methods, hydraulic methods, habitat quantification methods, and global

42 analysis methods (Dong et al., 2017; Sajedipour et al., 2017). The water level is an important index

43 to measure hydrologic regimes and ecosystem health of lake wetland (Wu et al., 2017; Zhang et al.,

44 2018b; Wu et al., 2020). Therefore, studies on EWRs of lake wetland mainly focus on ecological

45 water level (Ye et al., 2017; Wang et al., 2020b; Zhang et al., 2016). The early studies primarily

46 focused on the definition and research of the minimum ecological water level and lacked the
47 understanding of the relationship between water level and ecosystems, which was relatively simple

48 (Gan et al., 2015; Wang et al., 2020b). For example, the natural water level data method, ecological

49 water level method, lake morphology analysis method, minimum annual average water level method,

50 and minimum spatial demand method (Liang et al., 2013; Shang, 2013; Yang et al., 2019).

51 Additionally, Gan et al. (2015) suggested that a single characteristic value of water level cannot

52 maintain the ecological health of the lake wetland, and it is necessary to study the water level

53 regimes to maintain the structure, function, and process integrity of the lake wetland ecosystem,

54 including the monthly ecological water level threshold, the duration of high and low water level,

55 and the rate of water level change. However, it lacks a direct link with ecological indicators.

56 Furthermore, Zhang et al. (2016) calculated the minimum protection water level with waterfowl

57 richness as the target based on Response Surface Methodology (RSM). In general, the methods of

58 EWRs for wetland, which are based on the mechanistic relationships between hydrologic regimes

59 and ecological indicators, are current state-of-the-art approaches (Quadroni et al., 2017; Dong et al.,

60 2017).

61 Dongting Lake, the second-largest freshwater lake in China, located downstream of the TGD

62 on the Yangtze River, is a wintering habitat for migratory birds along the East Asian-Australasian

63 Flyway. East Dongting Lake, a major part of Dongting Lake, with giant areas of mudflats and moist

64 meadow can provide abundant food resource for migratory birds (Zhang et al., 2016; Hu et al., 2018).

65 In particular, the wetland attracts approximately 80% of migratory birds of Dongting Lake (Zhang

66 et al., 2018b; Zou et al., 2018). However, the TGD, a tremendous and comprehensive hydropower

67 project, has significantly changed the hydrologic regimes and had a great impact on habitat

68 suitability for migratory birds of East Dongting Lake (Wu et al., 2021; Yu et al., 2018). Previous
69 researches primarily focused on the alteration of TGD on hydrologic regimes, landscape patterns,

70 and migratory waterbird population distributions (Wu et al., 2017; Zou et al., 2018; Yu et al., 2018).

71 Moreover, numerous studies have been conducted on ecological water level based hydrologic data

72 (Liang et al., 2013; Shang, 2013; Wang et al., 2020b). However, the ecological water level of East

73 Dongting Lake wetland based on the mechanistic relationships between the water level and

74 ecological indicators remains poorly characterized.

75 In this study, we established a habitat suitability model of migratory birds feeding based on

76 Digital Elevation Model (DEM) and Normalized Difference Vegetation Index (NDVI). We aimed

77 to (1) calculate the ecological water level of East Dongting Lake in dry seasons based on the

78 mechanistic relationships between the water level and habitat suitability of migratory birds feeding,

79 (2) identify the influence of the TGD on habitat suitability of migratory birds feeding, (3) recognize

80 the change of habitat suitability of migratory birds feeding by comparing the transition of land-cover

81 in similar hydrologic years before and after the operation of TGD, and (4) reveal the driving factors

82 of wetland change and provide countermeasures for habitat conservation of East Dongting Lake.

83 2. Study area and dataset

84 2.1. Study area

85 Dongting Lake, the second-largest freshwater lake in China, is located downstream of the TGD

86 on the Yangtze River, including East Dongting Lake, West Dongting Lake, and South Dongting

87 Lake. The water enters the lake by Four Rivers (Xiang, Zi, Yuang, and Li) and Three Outlets (Songzi,

88 Taiping, and Ouchi) and out of the lake by Chenglingji (Fig. 1).

89 East Dongting Lake, a major part of the Dongting Lake, is a vital wintering habitat for

90 migratory birds along the East Asian-Australasian Flyway. The annual average air temperature is

91 17.4 °C (1959-2020). The annual average precipitation is 1327 mm (1959-2020). Precipitation from
 92 April to September accounted for more than 60% of the total annual precipitation due to the

93 influence of subtropical monsoon climate (Yang et al., 2020). The water level began to decline

94 rapidly in October, and there are giant exposed lake bottomlands, which can provide a wintering

95 habitat for waterfowl (Zhang et al., 2016). Every year, about 80% of the migratory birds of Dongting

96 Lake come to the East Dongting Lake to overwinter (Zou et al., 2018). In addition, the Carex

97 community, the main food resource of migratory birds, is the predominant vegetation in East

98 Dongting Lake (Zhang et al., 2018b).

99 The TGD, a tremendous and comprehensive hydropower project, is located on the upper of the

100 Dongting Lake on Yangtze River (Fig. 1). The TGD, which began operation in 2003, stored water

101 to 175 m for the first time in September 2010. More specifically, the water level should lower to

102 145 m in June for the flood control and water was required to release from January to March for the

103 water supply.

104

105 Fig. 1. Locations of the East Dongting Lake, Chenglingji station, and the TGD in China.
106 2.2. Datasets

107 In this study, long-term remote sensing (Landsat) data and DEM data were collected from the

108 United States Geological Survey (http://www.usgs.gov/). The Landsat images are shown in Table 1.

109 The DEM data is a 30-m spatial resolution (2009). The hydrological data (water level, runoff,

110 sediment) were collected from the Institute of Water Conservancy and Hydroelectric Science of

111 China. The air temperature and precipitation data were collected from the China Meteorological

112 Data Sharing Service System. The migratory bird data was collected from East Dongting Lake

113 National Nature Reserve.

114 Table 1 Landsat images used in this study. Hydrologic year (HY) starts on 1st April and ends on

115 31st March. Landsat-4: 1989/02/11, Landsat-8: 2013-2020, others are Landsat-5 images.

HY Date HY Date HY Date HY Date

1987 1987/12/31 1995 1995/12/05 2003 2004/02/13 2013 2014/01/23
1989/02/11 1996 1996/12/07 2006/11/01 2014 2015/03/31
1988 2006
1989/03/07 1997 1998/02/12 2006/12/19 2015/11/26
2015
1989 1989/12/04 1999 1999/12/24 2008/12/08 2016/03/01
1991 1991/11/08 2000 2001/01/11 2008 2008/12/24 2016/11/28
2016
1992 1993/01/29 2001 2001/12/25 2009/03/14 2017/02/16
1993/11/13 2001 2002/01/22 2010/11/12 2017/12/17
1993 2010 2017
1993/12/31 2002 2003/01/17 2011/03/04 2018/02/03
1994 1995/0204 2003 2003/12/27 2013 2013/12/06 2020 2020/12/25

116

117 3. Methods
118 3.1. Data pre-processing

119 Landsat images should be pre-processing, which can minimize the influence of sensor, solar,

120 atmosphere, and topography. The pre-processing steps of Landsat images are very common, mainly

121 including radiometric calibration and atmospheric corrections. Those are done through the ENVI

122 5.3 software, and a more detailed image pre-processing can be found in Yang et al. (2020).
123 3.2. Classification method for Landsat images

124 The classification of the East Dongting Lake’s land-cover is various for meeting different

125 research purposes. According to previous studies (Chen et al., 2016; Wu et al., 2021), the land-cover

126 of shallow water, Mudflat, Tender Marsh land (TML), and Marsh land (ML) is considered suitable

127 for migratory birds, whereas the land-cover of deep water and Reed beach (RB) is not suitable for

128 migratory birds. In this study, there are five habitat types (Water, TML, ML, Mudflat, RB) in East

129 Dongting lake wetland, which were classified by the Support Vector Machine (SVM) approach. The

130 SVM method, a common supervised classification method, was found by Cortes and Vapnik (1995).

131 Additionally, high-resolution images were used for accuracy verification, which was obtained from

132 Google Earth™ (http://earth.google.com), and the overall accuracy was satisfied.

133 In this study, the submerged conditions can directly affect the lake bottom’s exposure time and

134 distribution of vegetation at the junction of land and water (Zhou et al., 2020; Yang et al., 2020).

135 Because of the comprehensive action of water level and topography, the soil moisture is higher as

136 the distance from the lake is closer, and gradually decreases as the elevation increases slowly and

137 the distance from the lake is further (Wu et al., 2020; Zhou et al., 2020). The difference in soil

138 moisture at different elevations is a key dominant factor influencing vegetation distribution, which

139 had a great impact on habitats suitability for migratory birds feeding (Wu et al., 2020; Zhou et al.,

140 2020; Yang et al., 2020). Therefore, land-cover and DEM are considered important factors affecting

141 habitat suitability. In this study, NDVI, the red and NIR ratio, was used to extract the desired land-

142 cover (Fig. 2), as follows:

NIR  R
143 NDVI  (1)
NIR  R
144 The NIR and RED band of different Landsat images can be found in Yang et al. (2020).
145 3.3. Habitat suitability model of migratory birds feeding based on DEM and NDVI

146 In this study, the WUA of deep water and RB is equal to 0, which is considered not suitable for

147 migratory birds feeding. The WUA of shallow water areas, RB, TML, and ML depends on DEM,

148 which is considered suitable for migratory birds feeding. As shown in Fig. 2, different land-cover

149 was classified by NDVI values, and the classification values of Landsat images in different periods

150 were different. Moreover, the habitat suitability criteria curve of DEM for migratory birds feeding

151 is divided as follows, which is based on the different land-cover on different DEM. When DEM is

152 less than or equal to 24 m, the suitability is equal to 1 (A1=1). When DEM is equal to 25 m, the

153 suitability is equal to 0.9. When DEM is equal to 26 m, the suitability is equal to 0.8. When DEM

154 is equal to 27 m, the suitability is equal to 0.7. When DEM is equal to 28 m, the suitability is equal

155 to 0.6. When DEM is larger than 28 m, most of the habitat types are RB, which is not suitable for

156 migratory birds feeding.

157

158 Fig. 2. Habitat suitability model of migratory birds feeding based on DEM and NDVI.

159 4. Results and discussion

160 4.1. Ecological water level based on habitat suitability model

161 The responses of the WUA to water level were shown in Fig.3. As shown in formula (2), the

162 mechanistic relationships between the water level and WUA were evaluated quantificational by the
163 linear regression method.

23.704Z  14.4 Z  20.06


164 WUA  94.653Z  1437.7 20.06  Z  20.93 (2)
71.088Z  2031.6 Z  20.93


165 where, the WUA is the weighted usable area of migratory birds feeding (km2), and Z is the water

166 level of Chenglingji station (m).

167

168 Fig. 3. Responses of the WUA to the water level of East Dongting Lake in dry seasons.

169 As shown in Fig. 3, the WUA varies with the water level in a single peak pattern, reaching the

170 maximum when the water level is 20.93 m at Chenglingji Station, about 534.73 km2. The changing

171 trend of the WUA was divided into three different stages. Furthermore, the WUA decreased rapidly

172 and then slowly and finally stabilized at about 450 km2 when the water level is less than 20.06 m,

173 indicating that there is still a certain amount of area for migratory birds feeding when the lake water

174 level is below a certain threshold, which mainly refers to Mudflats. However, when the lake level

175 is greater than 23 m, the water will cover most of the Mudflats and Carex community, indicating

176 that there will be inadequate food resource and habitat for plentiful migratory birds (Zou et al., 2017;

177 Zhang et al., 2018b; Zou et al., 2018).

178 Only if there is enough suitable habitat in East Dongting Lake can plentiful migratory birds

179 feeding be guaranteed. In this study, according to Fig. 3, when WUA is greater than 500 km2, the
180 water level ranges from 20.47-21.55 m, which is considered the most suitable ecological water level

181 in East Dongting Lake. When WUA is greater than 450 km2, the water level ranges from 19.59-

182 22.25 m, which is considered a minimum and maximum ecological level.

183 Comparison of research results on the ecological water level in East Dongting Lake based on

184 different approaches were summarized in Table 2. The minimum ecological water level based on

185 the natural water level data method and ecological water level method, is the constraint on the

186 average annual minimum water level of the lake (Yang et al., 2019). The minimum ecological water

187 level based on level-area-storage curves and lake morphology analysis method is the water level

188 corresponding to the maximum change of lake area with the water level, which mainly protects the

189 lake ecosystem from the perspective of lake topography (Shang, 2013; Yang et al., 2019). The

190 calculation results of the above two methods cannot be directly compared with that of this study due

191 to different purposes. Liang et al. (2013) considered the monthly mean water level with the highest

192 probability as the ecological water level. Wang et al. (2020b) calculated the ecological water level

193 process from the whole ecosystem based on the IHA-RVA method. The results of the above two

194 methods are similar with those of this study. Zhang et al. (2016) used RSM to calculate the minimum

195 protection water level with migratory bird richness as the target. The results are larger than those in

196 this study because they are based on the populations of migratory birds in the whole Dongting Lake.

197 In general, the finding can provide a reference for habitat conservation of migratory birds in East

198 Dongting Lake.

199 Table 2 Comparison of research results on the ecological water level in East Dongting Lake.

Source of research Time series of The ecological water

Research methods
results water level level in dry seasons
Liang et al. (2013) 1952-2006 Four probability functions 19.13-20.85 m
Shang (2013) - Level-area-storage curves 26.7 m
 Zhang et al. (2016) 2005-2012 RSM 22.61–23.49 m
Annual guarantee rate method
Yang et al. (2019) 1953-2013 Ecological water level method 22.62 m
Lake morphology analysis method
Wang et al. (2020b) 1959-2016 IHA-RVA 19.96-21.27 m
In this study 1987-2020 Habitat suitability model 20.47-21.55 m

200 4.2. Influence of the TGD on WUA changes in dry seasons

201 As shown in Fig. 4a, among the alteration of average WUA in dry seasons, the most prominent

202 change occurred in January, which increased by 6.51%. Moreover, the average WUA increased by

203 2.41% in December and decreased by 5.27% in February after 2003. As shown in Figure 4b, the

204 average WUA is greater than 500 km2 and 450 km2 in dry seasons are advanced by 15 d and 19d,

205 respectively, after 2003. As shown in Figure 4c, the average number of days with WUA greater than

206 500 km2 increased by 23.21 d, and the number of days with WUA greater than 450 km2 increased

207 by 4.16 d after 2003.

208 According to Fig. 3, there is a linear correlation between the water level and WUA. Therefore,

209 alteration of the water level in dry seasons were analyzed. The results demonstrated that the average

210 minimum water level advanced 20 d (Fig. 4b) and rose by 0.66 m (Fig. 4c) in dry seasons after 2003.

211 The box plot showed that the water level of minimum value, P_ (25) value, median value, P_ (75)

212 value, maximum value increased obviously in January. The average water level rose by 0.77 m in

213 January and 0.65 m in February, and decreased by in December 0.23 m after 2003. The number of

214 migratory birds is maximum in January of East Dongting Lake (Zhang et al., 2016). The increase

215 of average WUA due to the operation of the TGD is beneficial to migratory birds feeding. Moreover,

216 what’s noteworthy is that the fluctuation of the lowest water level in the dry seasons is only 1.31m,

217 which accounts for 41.59 % in the pre-TGD period (Fig. 4c). The fluctuation of water level had an

218 important effect on the ecological environment of lake (Wu et al., 2021; Gan et al., 2015; Leira and
219 Cantonati, 2008). Further study is required to learn the influence of water level fluctuation on

220 migratory birds, fish, and vegetation in dry seasons of East Dongting Lake.

221 Fig. 4. Comparison between the pre- and post-TGD period, (a) the average WUA in December,

222 January, and February, (b) the average start date of WUA>500 km2, WUA>450 km2, and minimum

223 water level in dry seasons, (c) duration days of WUA>500 km2, WUA>450 km2 and minimum water

224 level in dry seasons, (d) box plot showing water level in December, January, and February.

225 4.3. Influence of the TGD on land-cover changes in dry seasons

226 As shown in Fig. 5a and Table 3, the TML expanded southward and increased from 308.74

227 km2 in 2001 to 589.28 km2 in 2013, which was the most significant change of dry hydrologic years.

228 The RB shrank southward and decreased from 391.53 km2 in 2001 to 208.98 km2 in 2013, which

229 was the second obvious change of dry hydrologic years. However, land-cover transition of mudflat,

230 water, and ML presented insignificant changes. The land-cover transition of TML and RB of normal

231 hydrologic years was similar to that during dry hydrologic years. TML increased from 155.37 km2

232 in 2000 to 524.39 km2 in 2015 and RB decreased from 365.86 km2 in 2000 to 141.96 km2 in 2015
233 (Fig. 5b and Table 3). However, the variation of TML and RB of flood hydrologic years was the

234 inverse of that during dry hydrologic years. TML decreased from 328.46 km2 in 1999 to 168.80 km2

235 in 2020 and RB increased from 174.75 km2 in 1999 to 278.91 km2 in 2020 (Fig. 5c and Table 3).

236 In general, the mutual transition of Mudflat, TML, ML, RB, and Water was gathered at the

237 junction of Water, ML, and TML. Moreover, the transition area shrank dramatically after the

238 operation of TGD, for example, the Mudflat area decreased from 143.47 km2 in 2001 to 70.74 km2

239 in 2014 during dry hydrologic years and from 144.14 km2 in 2000 to 74.79 km2 in 2015.

240 Furthermore, the most prominent transition among changes of land-cover was that the RB (155.07

241 km2) and ML (150.66 km2) changed to TML during dry hydrologic years, the RB (171.90 km2) and

242 ML (174.30 km2) changed to TML during normal years, which was similar that during dry

243 hydrologic years, and the ML (130.31 km2) changed to RB during flood hydrologic years. Although

244 the change of land-cover in East Dongting Lake was complicated after the operation of TGD, the

245 RB area, an unsuitable area for birds, decreased remarkably during dry and normal hydrologic years,

246 which was unquestionable.

247

248 Fig. 5. Land-cover transition of dry, normal, and flood hydrologic years in the East Dongting

249 Lake.

250 Table 3 Land-cover transition matrix of dry, normal, and flood hydrologic years in the East

251 Dongting Lake (unit: km2).

Dry hydrologic year

2001\2013 TML ML RB Mudflat Water Total
TML 187.52 75.57 21.07 9.13 15.45 308.74
ML 150.66 48.73 3.66 3.96 5.44 212.44
RB 155.07 38.65 181.72 3.16 12.78 391.38
Mudflat 74.02 8.78 1.00 34.79 24.88 143.46
Water 22.01 2.59 1.52 19.71 132.38 178.20
Total 589.28 174.31 208.97 70.74 190.92 1234.23
Normal hydrologic year
2000\2015 TML ML RB Mudflat Water Total
TML 81.83 174.30 171.90 60.00 36.75 524.77
ML 56.71 90.52 63.12 21.61 15.88 247.84
RB 2.81 32.32 102.26 2.05 2.93 142.36
Mudflat 4.60 3.26 7.64 36.12 23.15 74.78
Water 9.30 15.82 20.90 24.33 174.12 244.48
 Total 155.26 316.22 365.82 144.10 252.83 1234.23
Flood hydrologic year
1999\2020 TML ML RB Mudflat Water Total
TML 80.34 108.07 53.00 39.77 47.16 328.34
ML 50.66 225.78 130.31 7.41 34.27 448.43
RB 13.40 53.52 74.31 6.14 27.24 174.61
Mudflat 8.92 15.07 11.82 16.15 51.15 103.11
Water 15.88 19.25 9.38 37.53 97.70 179.74
Total 169.19 421.69 278.83 107.01 257.52 1234.23

252

253 4.4. The driving factors of habitat suitability change of migratory birds

254 The TGD, a tremendous and comprehensive hydropower project, has significantly changed the

255 intra- and inter-annual variation of hydrologic regimes and had a great impact on habitat suitability

256 of migratory birds in East Dongting Lake (Yu et al., 2018; Wang et al., 2020a; Zhang et al., 2018b;

257 Zou et al., 2018).

258 On the one hand, for inter-annual variation of hydrologic regimes, the amount of water

259 exchange in Dongting Lake altered slightly after 2003 (Fig. 6a). However, compared with the pre-

260 TGD period, the volume of water entering and out of Dongting Lake has been reduced by 305.90×

261 108 m3 and 313.28×108 m3, respectively. Compared with the pre-TGD period, the sediment discharge

262 of Three Outlets, Four River, and Chenglingji decreased by 90.35%, 63.83%, and 37.35%,

263 respectively (Fig. 6b). During the pre-TGD period, the sediment entering the lake from Three

264 Outlets accounted for 79.26% of the total sediment entering the lake. The TGD significantly reduced

265 the sediment entering the Dongting Lake from Three Outlets, which led to the transformation of the

266 Dongting Lake from siltation to scouring (Fig. 6b). The speed and mode of sedimentation played a

267 crucial role in the evolution trend and direction of lake wetland vegetation. The change from

268 deposition to erosion due to the operation of TGD directly affected the evolution trend and direction

269 of East Dongting Lake vegetation. For example, there is a negative correlation between deposition
270 and vegetation before 2003, but the TGD alleviated obviously sediment deposition and resulted in

271 the expansion of vegetation to the center of the lake (Yang et al., 2020). In addition, the operation

272 of TGD also changed transport patterns of sediments, which led to more clear water (low sediment

273 content) from Three Outlets to Dongting Lake (Yang et al., 2017; Li et al., 2018; Zhang et al., 2017).

274 These changes not only affected the evolution trend of wetland but also caused water quality

275 variations of East Dongting Lake (Geng et al., 2020, 2021).

276 On the other hand, for intra-annual variation of hydrologic regimes, the volume of water

277 flowing into Dongting Lake decreased dramatically from mid-September to late October because of

278 the water storage of TGD and the corresponding water level also declined (Fig. 6c). The average

279 water level of East Dongting Lake in September and October decreased by 1.46 and 1.89 m,

280 respectively (Fig. 6c), and the corresponding duration days of water level from 25 to 28 m decreased

281 by 17.15 d compared with the pre-TGD period (Fig. 6d), which advanced the dry seasons and further

282 exposed the bottomlands. However, with the dry seasons, it’s just the other way around. As a result

283 of the release of water from TGD, the mean water level of East Dongting Lake in January and

284 February increased by 0.77 m and 0.65 m, respectively (Fig. 6c), and the corresponding duration of

285 water level from 20 to 24 m increased by 43.14 d compared with pre-TGD period (Fig. 6d), which

286 extended the dry season and increased the area of water in dry seasons, indicating that the highest

287 elevation and mean topsoil moisture of bottomlands slightly increased after 2003 (Yang et al., 2020;

288 Zhou et al., 2020).

289 The submerged regimes of East Dongting Lake can be directly affected by the water level and

290 duration days, which had a great impact on habitat suitability of migratory birds (Zhang et al., 2020;

291 Wu et al., 2020). The change in submerged conditions due to the TGD directly affected the lake
292 bottom’s exposure time and distribution of vegetation at the junction of land and water (Yang et al.,

293 2020). In this study area, due to the comprehensive action of water level and topography, the soil

294 moisture is higher as the distance from the lake is closer, and gradually decreases as the elevation

295 increases slowly and the distance from the lake is further. The difference in soil moisture at different

296 elevations is a key dominant factor influencing vegetation distribution, which had a great impact on

297 habitats suitability of migratory birds feeding (Zhang et al., 2018b; Zou et al., 2018). The increase

298 of water level and duration days in dry seasons advanced and prolonged the lake bottom’s exposure

299 time, which resulted in the expansion of vegetation to the center of the lake (Wu et al., 2017; Hu et

300 al., 2018). Moreover, Zhou et al. (2020) find that water level fluctuation in wet seasons was the

301 determinant towards the distribution of Carex and Phragmites. When the average water level was

302 about 29 m in wet seasons, it will be the most suitable for the growth of Carex, which is the main

303 food resource of wintering birds. As shown in Fig. 6d, after 2003, the duration days of water level

304 from 28 to 30 m increased by 6.96 d, indicating that the water level during the post-TGD period is

305 more suitable for the growth of the main food resource of migratory birds. This may be an important

306 reason for the increase of migratory birds in East Dongting Lake (Fig. 7).
307 Fig. 6. Comparison between the pre- and post-TGD periods, (a) Water exchange, (b) Sediment

308 exchange, (c) Monthly mean water level, (d) Duration days of different water level, (e) Monthly

309 mean air temperature, and (f) Monthly mean local precipitation in East Dongting Lake.

310 What’s more, meteorological elements also played a crucial role in the evolution of wetland

311 vegetation (Yang et al., 2020). No significant change was observed in air temperature and

312 precipitation in dry seasons, indicating that wetland habitat suitability in East Dongting Lake was

313 limited affected by air temperature and precipitation in dry seasons (Fig. 6e and 6f). However, air

314 temperature rose from July to September inhibited the vegetation growth of East Dongting Lake far

315 from the center of the lake (Zhou et al., 2020). Drought can significantly reduce wetland habitat

316 suitability and affect the populations of migratory birds (Fig. 7). Moreover, the Chinese government

317 implemented ecological restoration projects (Return land to lake in 2012 and Return forest to

318 wetland), which are beneficial for improving habitat suitability of East Dongting Lake.
319

320 Fig.7. The populations of migratory birds in January from 2006 to 2020 at the East Dongting

321 Lake.

322 4.5. Implications for wetland conservation

323 East Dongting Lake is a significant wintering habitat for migratory birds. However, intense

324 human activities especially the operation of the TGD had a significant impact on the habitat of

325 migratory birds in East Dongting Lake, and climate change is not the main driving factor after 2003

326 (Zhang et al., 2016; Wu et al., 2021). Therefore, effective engineering measures should be put

327 forward to protect the habitat of migratory birds (Zhang et al., 2021). In this study, the most suitable

328 ecological water level can provide a reference for the protection of migratory birds in East Dongting

329 Lake. Moreover, more freshwater should be introduced through Three Outlets to improve the water

330 quality (Zhou and Zhang, 2018). Besides, the relevant law should be developed to protect wintering

331 habitat for migratory birds. Major land-cover change projects should be prohibited and the Poplar

332 Ecological Retreat project should be promoted, which can enhance ecological function and protect

333 wetland environment (Hu et al., 2020).

334 5. Conclusion
335 This study demonstrates a habitat suitability model to quantitatively analyze the effects of

336 water level change on the feeding area of migratory birds based on DEM and NDVI, and how does

337 the TGD affect the habitat suitability of migratory birds in East Dongting Lake. The major findings
338 demonstrated that the rise of water level in dry seasons and increase of duration days of water level

339 from 28 to 30 m in wet seasons due to the operation of TGD were the main reasons for the transition

340 of land-cover from RB and ML to TML during dry hydrologic years and normal hydrologic years,

341 which lead to the improvement of habitat suitability of migratory birds in East Dongting Lake.

342 However, severe drought can have significant adverse effects on the habitat suitability of migratory

343 birds. In addition, based on the habitat suitability model of migratory birds feeding, the most suitable

344 ecological water level in East Dongting Lake is 20.47-21.55 m. The findings can provide a useful

345 model for habitat conservation of migratory birds and a reference for the operation of TGD.

346 Acknowledgments:
347 This work was supported by the National Nature Science Foundation of China (Grant No.

348 51779094); Fund of Innovative Education Program for Graduate Students at North China University

349 of Water Resources and Electric Power, China (Grant No. YK2020-01).

350 CONFLICT OF INTEREST

351 The authors have no conflict of interest to declare.

352 References
353 Chen, Y., Niu, Z., Hu, S., Zhang, H., 2016. Dynamic monitoring of dongting lake wetland using time-series MODIS

354 imagery. J. Hydraulic Eng. 47, 1093-1104.

355 Cortes, C., Vapnik, V., 1995. Support-Vector Networks. Mach. Learn. 20, 273-297.

356 Dong, Z., Zhang, J., Zhao, J., 2017. Comments upon progress of environmental flows assessments. J. Hydraulic Eng.

357 48, 670-677.

358 Gan, F., Tang, L., Guo, H., Gao, W., 2015. New method and application of estimating ecological water level of the

359 Lake Poyang. J. Lake Sci. 27, 783-790.

360 Geng, M., Wang, K., Yang, N., Li, F., Zou, Y., Chen, X., Deng, Z., Xie, Y., 2020. Spatiotemporal water quality

361 variations and their relationship with hydrological conditions in Dongting Lake after the operation of the Three
362 Gorges Dam, China. J. Cleaner Prod. 283, 124644.

363 Geng, M., Wang, K., Yang, N., Li, F., Zou, Y., Chen, X., Deng, Z. Xie, Y., 2021. Evaluation and variation trends

364 analysis of water quality in response to water regime changes in a typical river-connected lake (Dongting

365 Lake), China. Environ. Pollut. 268, 115761.

366 Gleick, P. H., 1996. Basic Water Requirements for Human Activities: Meeting Basic Needs. Water Int. 21, 83-92.

367 Hu, J., Xie, Y., Tang, Y., Li, F., Zou, Y., 2018. Changes of Vegetation Distribution in the East Dongting Lake After

368 the Operation of the Three Gorges Dam, China. Front. Plant Sci. 09, 582.

369 Hu, W., Li, G., Gao, Z., Guan, J., Wang, Z., Li, Y., 2020. Assessment of the impact of the Poplar Ecological Retreat

370 Project on water conservation in the Dongting Lake wetland region using the InVEST model. Sci. Total

371 Environ. 733, 139423.

372 Leira, M., Cantonati, M., 2008. Effects of water-level fluctuations on lakes: an annotated bibliography.

373 Hydrobiologia. 613, 171-184.

374 Li, S., Yang, Y., Zhang, M., Sun, Z., Zhu, L., You, X., Li, K., 2018. Coarse and fine sediment transportation patterns

375 and causes downstream of the Three Gorges Dam. Front. Earth Sci. 12, 750-764.

376 Liang, J., Peng, Y., Guo, S., Li, X., Huang, L., Li, Z., Zeng, G., Xiao, Y., Shi, L., 2013. Determination of ecological

377 water-level in the eastern Lake Dongting with hydrological alterations. J. Lake Sci. 25, 330-334.

378 Prakasam, C., Saravanan, R., Kanwar, V. S., 2021. Evaluation of environmental flow requirement using wetted

379 perimeter method and GIS application for impact assessment. Ecol. Indic. 121, 107019.

380 Quadroni, S., Crosa, G., Gentili, G., Espa, P., 2017. Response of stream benthic macroinvertebrates to current water

381 management in Alpine catchments massively developed for hydropower Sci. Total Environ. 609, 484-496.

382 Sajedipour, S., Zarei, H., Oryan, S., 2017. Estimation of environmental water requirements via an ecological

383 approach: A case study of Bakhtegan Lake, Iran. Ecol. Eng. 100, 246-255.
384 Shang, S., 2013. Lake surface area method to define minimum ecological lake level from level-area-storage curves.

385 J. Arid Land. 5, 133-142.

386 Stone, R., 2008. Three Gorges Dam: into the unknown. Science. 321, 628-632.

387 Wang, H., Sun, F., Liu, W., 2020a. Characteristics of Streamflow in the Main stream of Changjiang River and the

388 Impact of the Three Gorges Dam. Catena. 189, 104498.

389 Wang, H., Zhu, Y., Zha, H., Guo, W., 2020b. Ecological water level and its safeguard measures in Lake Dongting. J.

390 Lake Sci. 32, 1529-1538.

391 Wang, J., Sheng, Y., Wada, Y., 2017. Little impact of the Three Gorges Dam on recent decadal lake decline across

392 China's Yangtze Plain. Water Resour. Res. 53, 3854-3877.

393 Wu, H., Dai, J., Sun, S., Du, C., Long, Y., Chen, H., Yu, G., Ye, S., Chen, J., 2021. Responses of habitat suitability

394 for migratory birds to increased water level during middle of dry season in the two largest freshwater lake

395 wetlands of China. Ecol. Indic. 121, 107065.

396 Wu, H., Zeng, G., Liang, J., Chen, J., Xu, J., Dai, J., Sang, L., Li, X., Ye, S., 2017. Responses of landscape pattern

397 of China's two largest freshwater lakes to early dry season after the impoundment of Three-Gorges Dam. Int.

398 J. Appl. Earth Obs. Geoinf. 56, 36-43.

399 Wu, H., Zheng, B., 2020. Wetland area identification and waterbird protection management in consideration of lake

400 topography and water level change. Glob. Ecol. Conserv. 23, e01056.

401 Yang, L., Wang, L., Yu, D., Yao, R., He, Q., Wang, S., Wang, L., 2020. Four decades of wetland changes in Dongting

402 Lake using Landsat observations during 1978-2018. J. Hydrol. 587, 124954.

403 Yang, Y., Du, C., Zhang, J., yU, G., Chen, H., Qian, Z., 2019. Determination of lowest ecological water level in

404 Dongting Lake area. Water Resour. Prot. 35, 93-98.

405 Yang, Y., Zhang, M., Li, S., Zhu, L., You, X., LI, k., Yu, X., 2017. Transport patterns of the coarse and fine sediments
406 and its causes in the downstream of the Three Gorges Dam. J. Lake Sci. 29, 942-954.

407 Ye, Z., Li, W., Chen, Y., Qiu, J., Aji, D., 2017. Investigation of the safety threshold of eco-environmental water

408 demands for the Bosten Lake wetlands, western China. Quat. Int. 440, 130-136.

409 Yu, Y., Mei, X., Dai, Z., Gao, J., Li, J., Wang, J., Lou, Y., 2018. Hydromorphological processes of Dongting Lake in

410 China between 1951 and 2014. J. Hydrol. 562, 254-266.

411 Zhang, C., Yuan, Y., Zeng, G., Liang, J., Guo, S., Huang, L., Hua, S., Wu, H., Zhu, Y., An, H., 2016. Influence of

412 hydrological regime and climatic factor on waterbird abundance in Dongting Lake Wetland, China:

413 Implications for biological conservation. Ecol. Eng. 90, 473-481.

414 Zhang, G., Wu, Y., Wu, Y., Liu, X., 2018a. A review of research on wetlands ecohydology. Adv. Water Sci. 29, 737-

415 749.

416 Zhang, H., Tang, W., Xin, X., Yin, W., 2021. Key strategies for the restoration of Dongting Lake in Middle Yangtze,

417 China. J. Environ. Sci. 100, 360-362.

418 Zhang, P., Zou, Y., Xie, Y., Zhang, H., Liu, X., Gao, D., Yi, F., 2018b. Shifts in distribution of herbivorous geese

419 relative to hydrological variation in East Dongting Lake wetland, China. Sci. Total Environ. 636, 30-38.

420 Zhang, P., Zou, Y., Xie, Y., Zhang, S., Chen, X., Li, F., Deng, Z., Zhang, H., Tu, W., 2020. Hydrology-driven

421 responses of herbivorous geese in relation to changes in food quantity and quality. Ecol. Evol. 10, 5281-5292.

422 Zhang, W., Yang, Y., Zhang, M., LI, Y., Zhu, L., You, X., Wang, D., Xu, J., 2017. Mechanisms of suspended sediment

423 restoration and bed level compensation in downstream reaches of the Three Gorges Projects(TGP). J. Geogr.

424 Sci. 27, 463-480.

425 Zhou, J., Wan, R., Wu, X., Zhang, Y., 2020. Patterns of long-term distribution of typical wetland vegetation (1987-

426 2016) and its response to hydrological processes in Lake Dongting. J. Lake Sci. 32, 1723-1735.

427 Zhou, J., Zhang, M., 2018. Effect of dams on the regime of the mid-lower Yangtze River runoff and countermeasures.
428 J. Lake Sci. 30, 1471-1488.

429 Zhou, Y., Ma, J., Zhang, Y., Li, J., Feng, L., Zhang, Y., Shi, K., Brookes, J. D., Jeppesen, E., 2019. Influence of the

430 three Gorges Reservoir on the shrinkage of China's two largest freshwater lakes. Glob. Planet. Change. 177,

431 45-55.

432 Zou, Y., Tang, Y., Xie, Y., Zhao, Q., Zhang, H., 2017. Response of herbivorous geese to wintering habitat changes:

433 conservation insights from long-term population monitoring in the East Dongting Lake, China. Reg. Envir.

434 Chang. 17, 879-888.

435 Zou, Y., Zhang, P., Zhang, S., Chen, X., Li, F., Deng, Z., Yang, S., Zhang, H., Li, F., Xie, Y., 2018. Crucial sites and

436 environmental variables for wintering migratory waterbird population distributions in the natural wetlands in

437 East Dongting Lake, China. Sci. Total Environ. 655, 147-157.